Review Article
Revision of Rubus (Rosaceae) in Taiwan
Juinn-Yih Huang(1) and Jer-Ming Hu(1*)
1. Institute of Ecology and Evolutionary Biology, National Taiwan University, 1, Roosevelt Rd., Sec. 4, Taipei 106, Taiwan.
＊ Corresponding author. Tel: +886-2-33662472; Fax: +886-2-23686750; Email: jmhu@ntu.edu.tw (Manuscript received 8 June 2009; accepted 3 August 2009)
ABSTRACT: The genus Rubus L. in Taiwan is taxonomically revised based on morphological and phytogeographical patterns. Forty taxa (34 species, 3 varieties, and 3 hybrids) of the genus are recognized, including two new hybrid species R. croceacanthus ×
corchorifolius and R. rosifolius × fraxinifolius. Among the Rubus taxa in Taiwan, R. croceacanthus H. Lév. exhibits tremendous
variation and we do believe that more investigations are needed in the future. Several taxa described in the Flora of Taiwan, 2nd ed. are treated as synonyms of other species, and several taxa in Taiwan are revised. The existence of R. howii Merr. & Chun, R.
linearifoliolus Hayata, and R. parvifraxinifolius Hayata in Taiwan are reconfirmed, we described R. howii fruit information which
was absent in original literature; and the third taxon is treated as a natural hydrid R. × parvifraxinifolius Hayata which putatively derived from R. rosifolius Sm. and R. linearifoliolus based on intermediate morphology and the scarcity of fruit set. Rubus
pseudoacer Makino ssp. flexuosus (Y.-C. Liu & F.-Y. Lu) H. Ohashi & C.-F. Hsieh is treated as synonym of R. pseudoacer Makino
and is excluded in Taiwan. A key to the taxa, synonyms and taxonomic descriptions of each taxon are presented. Moreover, information about phenology, chromosome number, distribution and specimens examined for each taxon are also provided. KEYWORDS: Rubus, Rosaceae, revision, hybrid, natural hybridization, Taiwan, taxonomy.
INTRODUCTION
Rubus L. (Rosaceae) is the only genus of Rubeae in
Rosoideae (Potter et al., 2007) and comprises about 700 species (Lu and Boufford, 2003). The genus is distributed worldwide and has been suggested to originate from Northern Temperate Zone, while only a few species are found in the tropics and/or in Southern Hemisphere (Kalkman, 2004).
There are over 40 Rubus taxa that have been described in Taiwan. Fifteen of the 40 Rubus taxa are endemic to Taiwan (37.5%), higher than the overall endemism of seed plants in Taiwan (26.2%) (Hsieh, 2003). Among the rest, 23 (57.5%) also occurred in Mainland China, 12 (30%) are also found in Japan, and 6 (15%) are also distributed in the Philippines.
Taxonomic history of Rubus in Taiwan could be traced back to 1896 by A. Henry's pioneer survey. Five
Rubus species, R. formosensis Kuntze, R. parvifolius L., R. rosifolius Sm., R. swinhoei Hance, and R. tagallus
Cham. & Schltdl., were recorded in A. Henry’s collecting list of Taiwan. The first comprehensive study on Rubus in Taiwan was done by Hayata in the 7th volume of Icones Plantarum Formosanarum published in 1918. He treated Rubus of Taiwan into 43 species and 1 variety and placed them in 3 subgenera, i.e.
Chamaebatus, Malachobatus, and Idaeobatus, following
the classification system proposed by Focke (1910, 1911, 1914). Detailed descriptions, leaf illustrations, and key to the species were all included in this master piece of work. Based on Hayata's framework, several taxonomic treatments of the genus Rubus in Taiwan have
been undertaken by Koidzumi (1930), Masamune (1954), Li (1963), Liu and Su (1977), Ying (1985), Hsieh (1989), Hsieh and Ohashi (1993), Lu et al. (2000), Naruhashi (2002), and Lu & Boufford (2003). However, opinions on species delimitation and taxonomic treatment varied and were at times inconsistent among researchers. The incongruences were likely due to the lack of comprehensive, broader geographical scale studies and also the frequent hybridization of Rubus species (Alice and Campbell, 1999), for example, the hybrids R.
croceacanthus × corchorifolius and R. rosifolius × fraxinifolius treated here.
The present work is based on newly collected materials, dried specimens, and related publications from neighboring regions of Taiwan, including Japan, China, and Malaysia. Voucher specimens examined were from the following herbaria: HAST, K, KYO, P, PE, PPI, TAI, TAIF, TCF, TI, TNM, and TNS. In this revision, 41 taxa (including 34 species, 4 varieties, and 3 hybrids) within the genus Rubus of Taiwan were treated. This study combined results from recent treatments on the genus Rubus (Lu et al., 2000; Naruhashi et al., 2002; Lu and Boufford, 2003; Hsu et al., 2007) and made many justifications from the Flora of Taiwan, 2nd ed. (Hsieh and Ohashi, 1993). Two new hybrids R. croceacanthus × corchorifolius and R.
rosifolius × fraxinifolius is added to the flora of Taiwan. Rubus parvifraxinifolius Hayata, which was previously
reduced to R. fraxinifoliolus Hayata (Hsieh, 1989), are now treated as a natural hybrid R. × parvifraxinifolius Hayata. The presence of R. howii Merr. & Chun and R.
linearifoliolus Hayata in Taiwan are reconfirmed, which
(1993). We add R. amphidasys Focke to the flora of Taiwan according to Hsu et al. (2007). We also made justifications on R. cardotii Koidz., R. croceacanthus H. Lév., R. fraxinifolius Poir., R. inopertus (Focke ex Diels) Focke, R. lambertianus Ser. var. morii (Hayata) S.-S. Ying, R. nagasawanus Koidz., R. nagasawanus Koidz. var. arachnoideus (Y.-C. Liu & F.-Y. Lu) S.-S. Ying, R.
parvifolius, R. pungens Cambess. var. oldhami (Miq.)
Maxim., R. reflexus Ker Gawl. var. hui (Diels ex Hu) F. P. Metcalf, R. rosifolius, R. rufus Focke, R. sumatranus Miq., R. swinhoei Hance var. kawakamii (Hayata) S.-C. Liu, R. taitoensis Hayata, and R. tephrodes Hance var.
setosissimus Hand.-Mazz. We re-elected R. incanus
Sasaki ex T.-S. Liu & T.-Y. Yang and R. pentalobus Hayata. Rubus pseudoacer Makino ssp. flexuosus Y.-C. Liu & F.-Y. Lu is treated as synonym of R. pseudoacer Makino and is excluded in the study.
Key to the taxa, synonyms for each taxon and taxonomic descriptions are presented. Moreover, information about phenology, chromosome number, distribution and specimens examined for each taxon are also provided.
TAXONOMIC TREATMENTS
Rubus L. 懸鉤子屬
Shrubs, usually scandant or prostrate, sometimes erect, only few species herbaceous; deciduous or evergreen; twigs and other parts nearly always with prickles. Leaves alternate; pinnately or palmately compound or simple, when simple usually lobed; stipules free or attached to the petiole base, persistent or caducous. Inflorescences terminal, determinate, elaborate, little or not branched, rarely reduced with solitary flowers. Flowers mostly bisexual, rarely unisexual and the plants dioecious; hypanthium saucer or cup shaped; sepals 5, imbricate, often unequal, connate near the base, exposed margins lobed or not; corolla 5, imbricate, mostly white, sometimes pink or purplish, in few species absent; stamens numerous, free; carpels numerous, free, on an elevated torus, each containing 2 ovules, usually only one developing, the styles filiform, nearly terminal, persistent. Fruits consisting of many 1-seeded drupelets, exocarp red, black, yellow or orange, mesocarp juicy, drupelets cohering and falling as a collective fruit without or together with the dry torus.
A large, complex genus of about 700 species, cosmopolitan in distribution, particularly abundant in the Northern Hemisphere; 40 taxa including 34 species, 3 varieties, and 3 hybrids in Taiwan.
Key to the species and varieties
1. Leaves compound; stipules linear or filiform, adnate to the basal part of petiole...2 1. Leaves simple; stipules linear or broad, adnate to the basal part of petiole, or at the junction of twig and petiole...17 2. Abaxial surface of leaflets densely whitish tomentose; flowers pink to purplish red. ...3 2. Abaxial surface of leaflets pubescent or glabrescent; flowers white or pink...5 3. Leaflets rounded or obtuse at apex; calyx abaxially prickled... ...19. R. parvifolius 3. Leaflets acuminate or acute at apex; calyx abaxially not prickled. ...4 4. Leaflets 3-7; inflorescences narrow cymose panicles...11. R. incanus 4. Leaflets 3; inflorescences corymbs. ...16. R. mesogaeus 5. Leaves 3-foliolate; stems and petioles with densely long, reddish
bristles; aggregate fruits yellow...35. R. wallichianus 5. Leaves 3-11-foliolate; stems and petioles pubescent, glabrous or with glandular hairs; aggregate fruits red, dark red, purplish black or orange. ...6 6. Flowers many in cymose panicles, or umbellate cymes. ...7 6. Flowers 1-several in terminal corymbs. ...9 7. Leaves 5- or rarely 3-foliolate, abaxially with dense hairs;
inflorescences narrow cymose panicles. ...18. R. parviaraliifolius 7. Leaves 7-11- or rarely 5-foliolate, glabrous; inflorescences broad lax cymose panicles, or umbellate cymes. ...8 8. Inflorescences umbellate cymes; flowers pink; leaves usually
9-11-foliolate. ... 12. R. inopertus 8. Inflorescences broad lax cymose panicles; flowers white; leaves
usually 7-9-foliolate...9. R. fraxinifolius 9. Plants dwarf, 5-10 cm tall; leaflets very small, 1-1.5 cm long, glabrous.
...32. R. taiwanicolus 9. Plants more than 10 cm tall; leaflets more than 1.5 cm long, glabrous or pubescent. ...10 10. Leaves 7-9(-11)-foliolate on sterile branch, occasionally 5-foliolate at base of inflorescence; plants glabrous, with sessile to stalked reddish glands, especially on twigs and pedicels... 14. R. linearifoliolus 10. Leaves 3-7(-9)-foliolate; plants glabrous or pubescent, with glandular hairs, glands, or not...11 11. Plants with reddish glandular hairs. ...12 11. Plants without glandular hairs, scarcely with sparse and short
glandular hairs. ...13 12. Flowers 2.5-5 cm in diam.; petals suborbicular, longer than sepals; aggregate fruits globose or oblong, red at maturity. ... ...6. R. croceacanthus 12. Flowers 1-2 cm in diam.; petals oblong to obovate, slightly shorter than sepals; aggregate fruits oblong, orange-red at maturity. ... ... 29. R. sumatranus 13. Plants glabrescent or glabrous; leaflets double-serrate or loosely
serrate...14 13. Plants pubescent, rarely glabrous; leaflets incised, double-serrate...15 14. Leaflets irregular- or loose-serrate, the terminal one
oblong-lanceolate; flowers 2.5-5 cm in diam...4. R. cardotii 14. Leaflets double-serrate, the terminal one ovate or elliptic; flowers 1-2 cm in diam. ...20. R. × parvifraxinifolius 15. Leaves, twigs and calyx pale yellow glandular-punctate, especially when young... 27. R. rosifolius 15. Leaves, twigs and calyx not glandular-punctate. ...16 16. Calyx with a few to many setose glandular hairs; twigs thinly
glandular-hairy; terminal leaflets ovate-lanceolate, cuneate at base . .. ...6. R. croceacanthus 16. Calyx with needle-like prickles and stalked glands; twigs pubescent or glabrescent; terminal leaflets ovate, triangular-ovate, rounded or subcordate at base. ... 23. R. pungens var. oldhamii 17. Stipules linear or filiform, adnate to the basal part of petiole,
17. Stipules broad, often dissected, at the junction of twig and petiole, fugacious or persistent. ... 20 18. Leaves abaxially usually pale or whitish; flowers 3 or more, in short racemes; calyx glabrous outside. ... 34. R. trianthus 18. Leaves abaxially not whitish; flower solitary, terminal on short
lateral twigs, or few appearing leaf-opposed; calyx finely pubescent outside. ... 19 19. Calyx not prickled outside. ... 5. R. corchorifolius 19. Calyx prickled outside. ...31. R. taitoensis 20. Terminal inflorescences broad lax cymose panicles. ... 21 20. Terminal inflorescences subracemes or simple racemes... 26 21. Leaves oblong or ovate-oblong, rounded at base; petals prominent shorter than sepals; carpels 5 or more, drupelets 1-3. ... ...24. R. pyrifolius 21. Leaves broadly ovate or suborbicular, cordate at base; petals longer than or slightly shorter than sepals; carpels numerous, druplets more than 10... 22 22. Leaves abaxially pilose or glabrous; aggregate fruits yellow or red. ... 23 22. Leaves abaxially densely gray tomentose; aggregate fruits purplish black. ... 24 23. Leaf blade and inflorescences thinly pilose...13a. R. lambertianus 23. Leaf blade nearly glabrous, inflorescences glandular-punctate or
with short glandular hairs...13b. R. lambertianus var. morii 24. Leaves shallowly 5-7-lobed on sterile twigs, apically acute or
acuminate. ...33. R. tephrodes var. setosissimus 24. Leaves deeply 5-lobed on sterile twigs, apically obtuse or acute. 25 25. Twigs with densely long glandular hairs and bristles. ... ... 17a. R. nagasawanus 25. Twigs without glandular hairs and bristles... ...17b. R. nagasawanus var. arachnoideus 26. Leaves oblong, ovate-oblong, or ovate, margin unlobed, undulate, or with shallow lobes on each side below middle. ... 27 26. Leaves suborbicular or broadly ovate, margin 3-5-lobed... 31 27. Leaves distinctly cordate at base; stipules divided deeply. ... 28 27. Leaves rounded, truncate, or shallowly cordate at base; stipules
apically laciniate. ... 29 28. Leaves oblong to ovate-oblong, with 1 or 2 broadly ovate shallow lobes on each side below middle. ... 10. R. howii 28. Leaves ovate or narrowly ovate, undivided or undulate... ... 7. R. flagelliflorus 29. Leaves abaxially yellowish brown tomentose; inflorescences
racemes... 15. R. liui 29. Leaves abaxially gray tomentose or glabrous; inflorescences
corymbose racemes. ... 30 30. Pedicels and calyx with purplish glandular hairs; leaves abaxially
gray tomentose. ... 30a. R. swinhoei 30. Pedicels and calyx with or without short stalked glands; leaves
abaxially glabrous. ... 30b. R. swinhoei var. kawakamii 31. Stipules and bracts laciniate shallowly to halfway, the lobes
lanceolate or ovate. ... 32 31. Stipules and bracts divided deeply, nearly to midvein or base, the
lobes linear, thread-like, or pinnately divided again into linear or linear-lanceolate lobules. ... 35 32. Leaves abaxially densely rusty yellow or rust colored... 33 32. Leaves abaxially densely gray or yellowish gray tomentose. ... 34 33. Leaves adaxially not distinctly bullate between veins; leaf lobes
acute or obtuse at apex; calyx lobes 6-8 mm long... ... 25. R. reflexus var. hui 33. Leaves adaxially distinctly bullate between veins; leaf lobes
rounded or obtuse at apex; calyx lobes 7-12 mm long ..26. R. rolfei 34. Stems prostrate, usually rooting at nodes; flowers solitary or 2-3 ... ... 22. R. pentalobus 34. Stems erect or scandent, not rooting at nodes; flowers several to
many, in subracemes...8. R. formosensis
35. Stems prostrate, usually rooting at nodes; leaves apically rounded or obtuse, rarely acute ………...……….. 36 35. Stems erect or scandent, not rooting at nodes; leaves apically acute to
caudate ……… 37 36. Leaves abaxially with needle-like prickles along veins, margin
3-lobed or undulate-lobed; inflorescences 1-2- flowered; calyx abaxially prickled ………....……… 21. R. pectinellus 36. Leaves abaxially unarmed, margin 3-5-lobed; inflorescences few
flowered, subracemes; calyx abaxially not prickled ….. 3. R. buergeri 37. Twigs with densely reddish brown long glandular hairs, usually
unarmed; stipule lobes not divided again …………. 2. R. amphidasys 37. Twigs without glandular hairs, prickled; stipule lobes pinnately
divided again into linear lobules……….…………. 38 38. Twigs with hairs, and sparse, recurved prickles; leaves adaxially
distinctly bullate between veins, lobes irregularly coarsely serrate; outer sepals palmately to pinnately laciniate .………. 1. R. alceifolius 38. Twigs with hairs, brownish bristles, and sparse, needle-like prickles;
leaves adaxially not distinctly bullate between veins, lobes irregularly sharply serrate; outer sepals apically shallowly laciniate... 28. R. rufus
1. Rubus alceifolius Poir. in Lamarck, Encycl. Meth. 6: 247. 1806; Hsieh, J. Taiwan Mus. 42: 104. 1989.
粗葉懸鉤子（羽萼懸鉤子）
Rubus moluccanus auct. non L.: Hayata, Icon. Pl. Formos. 1: 228. 1911.
Rubus laciniatostipulatus Hayata ex Koidz., J. Coll. Sci. Univ. Tokyo 34: 154. 1913.
Rubus pinnatisepalus auct. non Hemsl.: Li, J. Arnold. Arb. 25: 421. 1944.
Shrubs scandent. Twigs densely spreading straight tomentose-pilose, sparsely prickled. Leaves simple; blade suborbicular to broadly ovate, 7-20 × 6-18 cm, palmately 5-veined, abaxially yellowish gray colored tomentose, pilose along veins, adaxially sparsely pilose and distinctly bullate between veins, base cordate with basal incision to 4 cm, margin shallowly 5-7-lobed, lobes apically obtuse or acute, irregularly coarsely serrate; petiole 3-7 cm, densely pilose and prickled; stipules caducous, free, to 2 cm, deeply palmately or pinnately divided, tomentose abaxially, lobes filiform to linear, pinnately divided again into linear lobules. Inflorescences terminal, narrow cymose panicles or subracemes, 6-12 cm, or flowers few in axillary clusters; pedicels 5-10 mm, tomentose-pilose, with minute recurved prickles; bracts to 1.5 cm, pinnatifid to pinnatipartite, lobes filiform to linear. Flowers 1-1.6 cm in diam., hypanthium cup shaped; sepals ovate to triangular-ovate, 6-9 mm, abaxially tomentose-pilose, apex and margin of outer sepals laciniate, inner sepals entire, shortly pointed; petals white, suborbicular to elliptic, 6-9 × 4-8 mm, base clawed; stamens many, shorter than petals, filaments to 4 mm, anthers with few long hairs; carpels numerous, longer than stamens, ovary and style glabrous. Aggregate fruit red at maturity, globose, 1-1.8 cm in diam. Flowering season August to Octobor; fruiting season Octobor to December. 2n = 42.
Distributed in Southern China, the Philippines, Vietnam, Laos, Burma, Thailand, Cambodia, Malaysia,
and Indonesia. Taiwan, occurred in forests edges in the northern and central parts of the island at low altitudes 20-300 m.
Specimens examined: TAIWAN: Taipei City: Academia Sinica, T. I. Chuang 4642 (HAST); Nankang, J. M. Chao et al. 4841 (HAST); Hsinlung Rd., S. Y. Lu s.n. 19 Dec. 1972 (TAIF). Taipei Co.: Hsintien, Hsinwu Rd., W. C. Leong et al. 1264 (HAST); Ankeng, S. Sasaki s.n. 24 Nov. 1929 (TAI); Hsintien, Y. Shimada 11177 (TAI); Wulai, M. T. Kao s.n. 6 Aug. 1957 (TAI); Chuchih, T. Suzuki, 5421 (TAI); Kueishan, Masamune 2628 (TAI); Tachienshan, S. C. Liu et al. 718 (HAST); Sanhsia, S. Sasaki s.n. 13 Nov. 1910 (TAI). Ilan Co.: Chiaochi, C. M. Kuo 5689 (TAI); Toucheng, Huomanliaoshan, C. H. Lin et al. 861 (HAST); Yuanshan, Hsiatsukeng, J. J. Chen et al. 465 (HAST). Taoyuan Co.: Taikei, Y. Shimada 861A (HAST). Hsinchu Co.: Kuanhsi, Yushan stone area, S. Z. Yang 5241 (PPI); Peipu, S. Y. Lu 16884 (TAIF); Hengshan, Yulo, M. T. Kao, 7432 (TAI). Miaoli Co.: Tunglo, changshulin, C. H. Yu 128 (HAST); Tungho, Y. Shimada 869C (TAI); Toufen, Hsinglung Rd., T. Y. A. Yang et al. 11353 (TNM). Taichung Co.: Tungshih, H. Shimizu 540 (TAI); Hsinshe, Yuehhu Village, C. F. Chen 2781 (TNM); Tungmaoshan, G. P. Hsieh 2773 (TNM). Nantou Co.: Lienhuachih, W. P. Leu et al. 295 (HAST); Tiaumikeng, S. Y. Lu 16764 (TAIF); Puli, Liyutan, J. J. Chen et al. 438 (HAST); Sun Moon Lake, Kudo et al. 15508 (TAI). Changhua Co.: Kushan Temple, S. F. Huang 4941 (TAI); Shetou, Houtanching Trail, C. M. Wang et al. 11258 (TNM). Chiayi Co.: Meishan to Jueili, S. Y. Lu 16810 (TAIF).
2. Rubus amphidasys Focke, Bot. Jahrb. Syst. 29: 396. 1900; Hsu, Chiang & Peng, Taiwania 52: 113. f. 1.
2007. 周毛懸鉤子
Rubus rufus auct. non Focke: Naruhashi, Bot. Bull. Acad. Sin. 43: 194. 2002. pro parte.
Shrubs trailing or scandent. Twigs with dense reddish brown long glandular hairs, soft bristles, and long yellowish hairs, usually unarmed. Leaves simple; blade broadly to narrowly ovate, 5-11 × 3.5-9 cm, palmately 5-veined, both surfaces pilose, base cordate, margin 3-5-lobed, terminal lobe much larger and longer than lateral lobes, apex acute, lateral lobes obtuse, irregularly sharply serrate; petiole 3-5.5 cm; stipules free, to 1.5 cm, deeply palmately divided, lobes lanceolate to linear, with long glandular hairs, pilose. Inflorescences terminal, subracemes, 6-12 cm, 5-15-flowered, or in clusters of 3-5 in leaf axils; pedicel 5-12 mm, rachis, pedicels, and abaxial surface of calyx with dense reddish brown long glandular hairs, soft bristles, and yellowish hairs; bracts to 9 mm, lobed, lobes linear to lanceolate. Flowers 1-1.5 cm in diam., hypanthium cup shaped; sepals narrowly lanceolate, 1-1.5 cm, apex caudate, outer sepals usually 2- or 3-laciniate; petals white, broadly ovate to oblong, 4-7 × 3-5 mm, base shortly clawed, margin erose; stamens many; carpels many, glabrous, much longer than stamens. Aggregate fruit dark red at maturity, compressed globose, ca. 1 cm in diam., glabrous. Flowering season May to July; fruiting season July to September. 2n = 28.
Distributed in Southern China. Taiwan, occurred among shrubby thickets in the northern part of the island at low altitudes 300-700 m.
Specimens examined: TAIWAN: Taipei City: Waishuanghsi, abandonded Tsuishan Shooting Range, Chen et al. 448 (HAST); Taluntoushan, H. L. Chiang 468 (TAIF); Neihu, Pishan Camping Area, J. Y. Huang 2308 (HAST). Taipei Co.: Hsichih, Hsiwan Rd., Chen et al. 445 (HAST); Hsichih, Chepingliao, J. Y. Huang 194 (HAST); Wuchihshan, S. Y. Lu 14826 (TAIF).
Note: The chromosome counts were based on the description of ‘Rubus rufus’ in (Naruhashi et al., 2002). As Hsu et al. (2007) mentioned, the voucher specimen for R.
rufus [2n = 28: J. J. Chen et al. 445 (HAST)] Naruhashi et
al. (2002) cited should belong to R. amphidasys.
3. Rubus buergeri Miq., Ann. Mus. Bot. Lugduno-Batavi 3: 36. 1867; Li, Woody Fl. Taiwan 307. 1963. 寒莓
Rubus shimadai Hayata, Icon. Pl. Formos. 3: 94. 1913. Rubus pseudobuergeri Sasaki, Trans. Nat. Hist. Soc. Formos. 21: 249. 1931. - Rubus buergeri Miq. var. pseudobuergeri (Sasaki) T.-S. Liu & T.-Y. Yang, Sci. Ann. Taiwan Mus. 12: 7. 1969.
Subshrubs prostrate to erect. Stems slender, sometimes rooting at nodes, densely tomentose-pilose, unarmed or with sparse, minute prickles. Leaves simple; blade suborbicular to ovate, 4-15 cm in diam., palmately 5-veined, raised abaxially, both surfaces with hairs along veins, abaxially densely hirsute, adaxially slightly pubescent, base cordate to deeply cordate, margin shallowly 5-7-lobed, lobes apically rounded or obtuse, irregularly sharply serrate or dentate; petiole 4-8 cm, densely pubescent, sometimes with sparse prickles; stipules caducous, free, 8-15 mm, deeply palmately divided, lobes linear to linear-lanceolate, soft hairy. Inflorescences terminal, short subracemes, 4-5 cm, few flowered, or flowers several in axillary clusters; pedicel 5-10 mm, densely tomentose-pilose, with sparse prickles or not; bracts to 9 mm, similar to stipules. Flowers 6-10 mm in diam., hypanthium cup shaped, abaxially yellowish pilose and tomentose; sepals lanceolate or ovate-lanceolate, 5-9 mm, outer sepals often lobed apically, inner sepals entire, apex acuminate; petals white, obovate, 4-9 × 3-5 mm, apex erose; stamens many, shorter than petals, filaments glabrous; carpels glabrous, longer than stamens. Aggregate fruit red at maturity, globose, 0.6-1 cm in diam., glabrous. Flowering season August to October; fruiting season Octobor to January in next year. 2n = 56.
Distributed in Mainland China, Japan, and Korea. Taiwan, distributed in broad-leaved forests from low to medium altitudes 400-2000 m.
Specimens examined: TAIWAN: Taipei City: Tatungshan to Hsinpeitou, C. M. Kuo 9044 (TAI); Chihsingshan hiking trail, J. J. Chen et al. 450 (HAST), Hsiangtienchih, C. C. Liao 747 (HAST). Taipei Co.:
Szukanshui to Tatungshan, W. C. Leong et al. 1263 (HAST); Tataoshan, Y. C. Jeng 699 (TAI); Wulai, C. C. Chuang 2297 (TAI); Habun, T. Nakamura 605 (TAI); Mt. Peichatien, C. I Peng 7421 (HAST); Peichatienshan to Lupingshan, C. I Huang et al. 1387 (HAST); Roppeizan, T. Suzuki 18946 (TAI); Takaiyama, T. Suzuki 18524 (TAI). Ilan Co.: Ayushan, Y. C. Kao et al. 624 (HAST); Fushan Botanical Garden, H. L. Chiang 103 (HAST, TAIF); Mt. Fanpao, M. W. Jian et al. 106 (HAST); Taipinshan, S. F. Huang 5000 (TAI). Taoyuan Co.: Takuanshan Forest Reserve, C. I Peng et al. 14634 (HAST). Hsinchu Co.: Wufeng, Tienhu, C. M. Wang 10969 (TNM); Yufeng Village, en route from Szumakuszu to Sacred Tree, C. C. Liao 886 (HAST); Yuanyang Lake, J. J. Chen et al. 387, 390 (HAST). Miaoli Co.: Mt. Malapan, P. F. Lu 9850 (TAIF). Taichung Co.: Low Altitute Experimental Station to the hiking entrance of Sanchiaoshan, H. Y. Shen et al. 250 (HAST). Nantou Co.: Hsini to Hoshe, W. P. Leu et al. 1079 (HAST); Jenai, Tatung Village, H. F. Yen et al. 78 (TNM). Chiayi Co.: Chuchi, Shihcho, H. F. Yen 2930 (TNM). Kaohsiung Co.: Meilan logging track 14-15 km, C. I Peng et al. 18136 (HAST). Pingtung Co.: Chingshuiyin, H. Shimizu 3634 (TAI); Lilungshan, Y. K. Chen et al. 812 (HAST). Hualien Co.: Chingshuishan, Shakatang logging track, C. C. Liao 462 (HAST); Yuli, Chinshui Farm, C. H. Lin et al. 467 (HAST). Taitung Co.: Tajen, K. P. Lo 818 (PPI).
4. Rubus cardotii Koidz., Fl. Symb. Orient-Asiat. 62. 1930. - Rubus rosaefolius Sm. var. formosanus Cardot in Lecomte, Not. Syst. 3: 306. 1916.
紅狹葉懸鉤子（梣葉懸鉤子）
Rubus fraxinifoliolus Hayata, Icon. Pl. Formos. 5: 46. 1915, non Poir. 1806 ‘fraxinifolius’. syn. nov.
Rubus rubro-angustifolius Sasaki, Trans. Nat. Hist. Soc. Formos. 21: 221. 1931. syn. nov.
Rubus croceacanthus H. Lév. var. glaber auct. non Koidz.: Hsieh & Ohashi in Huang et al., Fl. Taiwan 2nd ed. 3: 126.
1993. pro parte. - Rubus asper Wall. ex D. Don. var. glaber (Koidz.) C.-F. Hsieh, J. Taiwan Mus. 42: 106. 1989. pro parte.
Shrubs suberect or scandent. Twigs reddish, glabrous, sparsely prickled, prickles 2-5 mm long. Leaves imparipinnate, mostly 5-foliolate, sometimes 3-foliolate on flower bearing twigs; blade of leaflets oblong-lanceolate to ovate-lanceolate, lateral veins 7-9 per side, raised abaxially and impressed adaxially, terminal leaflet usually longer and wider than lateral leaflets, 4-9 × 1-3 cm, lateral leaflets often slightly oblique, 1.5-5 × 0.6-2.5 cm, both surfaces glabrous, abaxially usually reddish when young, with compressed brown glands near margin, usually with sparse, minute prickles along midvein, base rounded to truncate, margin irregularly or loosely serrate, apex acuminate to acute; petiole 3-5 cm, petiolule adaxially grooved and sparsely hairy, with sparse, minute prickles; stipules attached to the petiole base, linear to linear-lanceolate, 5-10 mm, glabrous. Inflorescences terminal, 1- or rarely 2-flowered; pedicel 2-2.5 cm, glabrous, with few compressed brown glands. Flowers 2.5-5 cm in diam., hypanthium saucer shaped, abaxially with few compressed brown glands; sepals oblong-triangular, 1-1.3 cm, glabrous and with few sessile glands, margin tomentose, apex caudate; petals white, obovate, 1.2-2 ×
1-1.5 cm, base clawed. Stamens many, shorter than petals; carpels numerous, style and ovary glabrous, torus shortly stalked. Aggregate fruit red at maturity, ellipsoid to globose, 1.5-2 cm in diam., glabrous. Flowering season March to May; fruiting season May to July. 2n = 14.
Endemic to Taiwan. Occurred in thickets from low to medium altitudes 100-2800 m.
Specimens examined: TAIWAN: Hsinchu Co.: Chutung, Shangpingkou, S. F. Huang 4590 (TAI); Mamei Rd. to Litung Lodge, Y. Y. Huang et al. 446 (HAST); Chenghsipao, S. C. Wu et al. 1827 (TAI); Tuchang to Kuanwu, C. I Peng et al. 14864 (HAST). Miaoli Co.: Shihtan, Tali Coal Mine, C. I Peng 15067 (HAST); Nanchuang, Tanan logging track, P. F. Lu 9428 (HAST). Taichung Co.: Tahsuehshan Forest Recreation Area, J. C. Wang et al. 12140 (HAST); Yunling Lodge, T. Y. Liu et al. 1026 (HAST). Nantou Co.: Mayfeng farm, W. P. Leu et al. 462 (HAST); Juiyenhsi, J. J. Chen et al. 422 (HAST); Prov. #14A Hwy 19.5 km, W. C. Leong et al. 1312 (HAST); Yunhai, S. F. Huang 1192 (TAI); Nenkao pass, C. F. Chen 1382 (PPI); Pilushanhsi, Prov. #8 Hwy 99.8 km, C. H. Lin et al. 505 (HAST). Chiayi Co.: Alishan, Prov. #18 Hwy 23-23.5 km, H. L. Ho et al. 1132 (HAST). Kaohsiung Co.: Taoyuan, Prov. #20 Hwy 133-134 km, C. C. Wang et al. 1061 (HAST). Hualien Co.: Pilu, C. M. Wang s.n. 29 Jul. 1992 (TNM); Mukuashan, T. Shimizu 4143 (TAI); M. T. Kao 4143 (HAST), 4487 (HAST); Wanjung logging track to Chitsaihu, S. H. Wu et al. 1364 (HAST).
Note: International Code of Botanical Nomenclature 53.3: When two or more generic or specific names based on different types are so similar that they are likely to be confused (because they are applied to related taxa or for any other reason) they are to be treated as homonyms. According to this rule, the name R. fraxinifoliolus is so easily confused with R. fraxinifolius, so we used the name
R. cardotii to instead of R. fraxinifoliolus.
From the original description and the specimens cited by Liu and Su (1977), Rubus rubro-angustifolius is different from R. fraxinifoliolus (treated as R. cardotii here) only by its narrower terminal leaflets; but the shape of terminal leaflets in the latter often varies even on the same individual according to our examination. Both of them have glabrous twigs, abaxially reddish young leaves, loosely serrate leaf margins, and compressed brown glands especially on pedicels, hypanthia, and abaxial surface near margin of leaves; therefore, we treat R.
rubro-angustifolius as a synonym of R. cardotii here. Although R. rubro-angustifolius have been placed under R. croceacanthus var. glaber (treated as synonym of R. croceacanthus here) by Hsieh and Ohashi (1993), R. cardotii is easily distinguishable from R. croceacanthus by
the characters described above.
5. Rubus corchorifolius L. f., Suppl. Pl. Syst. Veg. 263. 1781; Matsum. & Hayata, J. Coll. Sci. Univ. Tokyo 22:
120. 1906. 變葉懸鉤子
Rubus corchorifolius L. f. var. glaber Matsum., Bot. Mag. Tokyo 15: 157. 1901.
Rubus retusipetalus Hayata, J. Coll. Sci. Univ. Tokyo 30: 94. 1911. syn. nov.
Rubus shinkoensis Hayata, J. Coll. Sci. Univ. Tokyo 30: 95. 1911.
Rubus arisanensis Hayata, Icon. Pl. Formos. 3: 87. 1913. Rubus suishaensis Hayata, Icon. Pl. Formos. 7: 6. f. 5-b. 1918. Rubus arisanensis Hayata var. horishaensis Hayata, Icon. Pl. Formos. 7: 15. f. 12-c. 1918.
Shrubs suberect. Twigs prickled, finely hairy, glabrate in age. Leaves simple; blade variable in shape, ovate to ovate-oblong, 4-12 × 3-9 cm, abaxially densely finely pubescent when young, gradually glabrescent, with sparse, small prickles along midvein, adaxially finely pubescent along veins, base rounded to cordate, margin undivided or 3-5-lobed, usually 3-5-divided on sterile branches, irregularly sharply serrate to doubly serrate, apex acuminate; petiole 1.5-2 cm, with dense fine hairs when young, with sparse, minute prickles; stipules attached to the petiole base, linear-lanceolate or linear, to 5 mm, soft hairy. Inflorescences terminal on short lateral twigs, 1-flowered or few flowers appearing leaf-opposed; pedicel 1-1.3 cm, finely pubescent. Flowers 1.5-2 cm in diam., hypanthium campanulate; sepals ovate to triangular-ovate, 6-10 mm, abaxially densely finely pubescent, unarmed, apex acute to shortly acuminate; petals white, oblong or elliptic, 9-13 mm × 6-9 mm, longer than sepals, glabrous, base clawed, apex obtuse; stamens much shorter than petals, filaments glabrous; carpels many, slightly shorter than stamens, ovary pubescent. Aggregate fruit orange-red to red at maturity, ovoid-globose, 1-1.2 cm in diam., densely finely pubescent. Flowering season January to March; fruiting season March to May. 2n = 14.
Distributed in Mainland China, Japan, Korea, Vietnam, and Burma. Taiwan, commonly distributed from low to medium altitudes, 10-2000 m throughout the island.
Specimens examined: TAIWAN: Taipei City: Academia Sinica, C. I Peng 4477 (HAST); Waishuanghsi, Chungyang Community, K. F. Chung et al. 1129 (HAST); Tsutzuhu, W. P. Leu 880 (HAST). Taipei Co.: Sanhsia, Paichi, S. M. Liu et al. 604 (HAST); Pinglin to Shihting, C. I Peng 15380 (HAST); Shihting, Prov. #9 Hwy 25-30 km, S. C. Liu et al. 314 (HAST); Hsiping Rd., between Tienfengku and watchtower, Y. C. Kao et al. 273 (HAST); Wanli, Tachienchih, Y. Y. Huang et al. 286 (HAST); M. T. Kao 10252 (TAI). Ilan Co.: Taipinshan, S. Y. Lu 15669 (TAIF); Mingchih to Chishan, C. M. Wang 5462 (HAST); Sanhsin, T. C. Huang 10817 (TAI); Shuanglienpi, K. C. Yang 4628 (HAST). Taoyuan Co.: Lungtan, Kunlun Medicial Plant Garden, C. I Peng et al. 13688 (HAST); Paling to Lalashan, P. F. Chen 32 (TAI); Lalashan, S. C. Wu 2630 (HAST). Hsinchu Co.: Yuanyang Lake, T. Y. Liu 912 (HAST). Taichung Co.: Lileng logging track, C. M. Wang et al. 4759 (TNM); Tungmaoshan, K. C. Yang 2367 (TNM). Nantou Co.: Huisunlinchang, C. M. Wang s.n. 21 Jan. 1992 (TNM); Mayfeng farm, W. P. Leu et al. 435 (HAST). Chiayi Co.: Between Chushan peony garden and Chushan station of Forest Railway, T. Kawakami 954 (TAI). Hualien Co.: Wanjung logging track 47 km, Y. C. Kao et al. 490 (HAST). Taitung Co.: Hsinkanshan, T. Y. A. Yang et al. 4432 (TNM); Wulu logging track, Y. Y. Huang et al. 266 (HAST).
6. Rubus croceacanthus H. Lév., Repert. Spec. Nov. Regni Veg. 11: 33. 1912; Hsieh & H. Ohashi in Huang
et al., Fl. Taiwan 2nd ed. 3: 125. 1993. 虎婆刺
Rubus asper Wall. ex D. Don., Prodr. Fl. Nepal. 234. 1825, non J. Presl. & C. Presl. 1822; Koidz., J. Coll. Sci. Univ. Tokyo 34: 139. 1913.
Rubus tagallus auct. non Cham. & Schltdl.: Matsum., Bot. Mag. Tokyo 16: 4. 1902.
Rubus rosaefolius Sm. var. hirsutus Hayata, J. Coll. Sci. Univ. Tokyo 25: 81. 1908. - Rubus hirsutus (Hayata) Hayata, Icon. Pl. Formos. 1: 227. 1911, non Thunb. 1813. syn. nov.
Rubus asper Wall. ex D. Don. var. pekanius Focke, Biblioth. Bot. 17 (Heft 72): 159. 1911.
Rubus sorbifolius auct. non Maxim.: Hayata, J. Coll. Sci. Univ. Tokyo 30: 96. 1911.
Rubus piptopetalus Hayata ex Koidz., J. Coll. Sci. Univ. Tokyo 34: 141.1913.
Rubus dolichocephalus Hayata, Icon. Pl. Formos. 3: 92. 1913. syn. nov.
Rubus sphaerocephalus Hayata, Icon. Pl. Formos. 3: 94. 1913. Rubus glanduloso-calycinus Hayata, Icon. Pl. Formos. 5: 42. 1915. - Rubus croceacanthus H. Lév. var. glanduloso-calycinus (Hayata) S.-C. Liu in Lu et al., Trees Taiwan 1: 122. 2000. syn. nov.
Rubus euphlebophyllus Hayata, Icon. Pl. Formos. 5: 44. 1915. Rubus hirsutopungens Hayata, Icon. Pl. Formos. 5: 58. 1915. syn. nov.
Rubus parvipungens Hayata, Icon. Pl. Formos. 5: 65. 1915. syn. nov.
Rubus rosaefolius Sm. var. polyphyllarius Cardot in Lecomte, Not. Syst. 3: 306. 1916. - Rubus polyphyllarius (Cardot) Koidz., Fl. Symb. Or.-Asiat. 62. 1930. syn. nov.
Rubus somai Hayata, Icon. Pl. Formos. 7: 19. f. 14a. 1918. Rubus croceacanthus H. Lév. var. glaber Koidz., Fl. Symb. Or.-Asiat. 65. 1930. - Rubus asper Wall. ex D. Don. var. glaber (Koidz.) C.-F. Hsieh, J. Taiwan Mus. 42: 106. 1989. syn. nov. Rubus takasagoensis Koidz., Fl. Symb. Or.-Asiat. 68. 1930. syn. nov.
Rubus hayatanus Koidz., Fl. Symb. Or.-Asiat. 68. 1930. syn. nov.
Rubus hayatai Nemato ex Makino & Nemoto, Fl. Jap. 2nd ed.
514. 1931. syn. nov.
Rubus okamotoanus Koidz., Acta Phytotax. Geobot. 8: 110. 1939. syn. nov.
Rubus rosaefolius Sm. var. formosanus auct. non Cardot: Li, Woody Fl. Taiwan 314. 1963.
Rubus pungens auct. non Cambess.: Li, Woody Fl. Taiwan 317. 1963.
Rubus sumatranus auct. non Miq.: Hsieh, J. Taiwan Mus. 42: 118. 1989. pro parte.
Rubus pungens Cambess. var. oldhamii auct. non (Miq.) Maxim.: Hsieh & H. Ohashi in Huang et al., Fl. Taiwan 2nd ed. 3:
136. 1993.
Shrubs suberect or scandant. Twigs pubescent to glabrescent, with densely to sparsely glandular hairs, and sparse, straight to curved prickles to 4 mm. Leaves imparipinnate, usually 5-7(-9)-foliolate, sometimes 3-foliolate on flower bearing twigs; blade of leaflets variable in shape, terminal leaflet ovate-oblong to lanceolate, 1.5-7 × 0.7-5 cm, lateral leaflets ovate to lanceolate, 1-5 × 0.5-3 cm, both surfaces with dense to
sparse soft hairs along veins, or glabrescent, sometimes with glandular hairs, usually with some small prickles along midvein abaxially, base cuneate to rounded, lateral leaflets often oblique basally, margin doubly serrate or irregularly sharply serrate, apex acute to acuminate; petiole 1-5 cm, densely soft hairy to glabrescent, with intermixed glandular hairs and sparse, minute prickles; stipules attached to the petiole base, lanceolate to linear, to 10 mm, pubescent to glabrescent, with intermixed glandular hairs. Inflorescences terminal, corymbose, 1- to several flowered; pedicel 2-4 cm, with few small prickles, glandular hairs, soft hairy or glabrescent. Flowers 2.5-5 cm in diam., hypanthium saucer shaped; sepals triangular-oblong, 0.7-1.5 cm, margin entire, apex long caudate; petals white, suborbicular, 1-2 × 0.7-1.5 cm, abaxially shortly hairy; stamens many, shorter than petals; carpels numerous, ovary with sparse, soft hairs especially apically, usually with some stalked glands, torus shortly stalked. Aggregate fruit red at maturity, globose or ovoid, rarely oblong, 1-2 cm in diam., with few glandular hairs. Flowering season February to June; fruiting season April to August. 2n = 14.
Distributed in Japan, Korea, the Ryukyus, Vietnam, Laos, Burma, Thailand, Cambodia, and India. Taiwan, very common in open places from low to high altitudes, 10-3600 m throughout the island.
Specimens examined: TAIWAN: Taipei City: Chihsingshan, T. Suzuki 8307 (TAI); S. Sasaki s.n. 28 May 1932 (TAI); Sozan, Y. Shimada 860 (HAST); S. Suzuki 3288 (TAI). Taipei Co.: Pali, C. C. Hsu et al. 4392 (TAI); Tamsui to Suihsientou, T. Suzuki 8271 (TAI); Hsiaopingting, W. P. Leu 1660 (HAST); Wanli, Tachienchih, Y. Y. Huang et al. 285 (HAST); Hsichih, Wuchihshan Hwy, K. F. Chung et al. 1124 (HAST); Tapingting, M. T. Kao 10241 (TAI); Yinhotung, C. C. Hsu et al. 4315 (TAI); Kueishan to Huoshawchang, T. Suzuki 8372 (TAI); Wulai, H. L. Chiang 521 (TAIF). Ilan Co.: Pingyuan logging track 7 km, Y. C. Kao et al. 345 (HAST); Taipinshan, S. Sasaki s.n. 8 May 1932 (TAI); Ssuyuanyakou, C. I Peng et al. 17412 (HAST); Chialohu, C. I Huang et al. 817 (HAST); Fushan Botanical Garden, C. I Peng et al. 13876 (HAST). Taoyuan Co.: Nankan, Y. Kudo s.n. 5 May 1929 (TAI); Lungtan, Kunlun Medicial Plant Garden, C. I Peng et al. 13687 (HAST); Yangmei, Puhsin, W. C. Leong et al. 2727 (HAST); Fuhsing, Ishan village, C. C. Liao et al. 169 (HAST); Meikueichi, W. C. Leong et al. 2839 (HAST). Hsinchu Co.: Hengshan, Dashanbeishan, S. C. Wu 2420 (HAST); Kuanhsi, W. C. Leong et al. 1134 (HAST); Tuchang to Kuanwu, C. I Peng et al. 14862 (HAST); Mamei Rd., Litung Lodge, Y. Y. Huang et al. 447 (HAST); Taikang Rd., from Hsiuluan to Taikang, Y. Y. Huang et al. 876 (HAST); Chenhsipao, C. H. Lin et al. 285 (HAST); Yuanyang Lake, K. Y. Wang et al. 1062 (HAST). Miaoli Co.: Shihtan, Hsintien Village, C. I Peng et al. 15251 (HAST); Szumahsien to Tienkou, T. Y. Liu et al. 83 (HAST); Hsuehshan #230 logging track 13.5 km, S. H. Wu 1216 (HAST); Huoyenshan, H. L. Chiang s.n. 14 Jan. 1987 (HAST, TAIF). Taichung City: Takeng, S. T. Chiu et al. 3273 (HAST). Taichung Co.: Taiping, Toupienkeng, K. Y. Wang et al. 796 (HAST); Sonhou, M. T. Kao 9644 (TAI); Pahsienshan Forest Recreation Area, T. Y. Liu et al. 885 (HAST); Huanshan, Y. Y. Huang et al. 396 (HAST); Wuling Lodge, C. I Peng et al. 17196 (HAST); Yunling Lodge to Tuochiatunshan, W. C. Leong et al. 3252 (HAST); Hsueshan, G. Masamune 1256 (TAI); Nanhutashan, N. Hukuyama et al. 17747 (TAI). Nantou Co.: Tsaotun, Chiuchiufeng, C. I Peng et al. 19247 (HAST); Hueisun Exp. Forest, C.
C. Chen 629 (TAIF); Puli, Liyutan, C. I Peng et al. 17167A (HAST); Lienhuachih, S. Y. Lu 3557 (TAIF); Wushe to Chienchin, C. S. Feung et al. 453 (TAI); Nengkao, K. Mori et al. 107 (TAI); Peitungyenshan, C. M. Wang 5519 (HAST); Chunyang, C. M. Wang 2600 (HAST); Mayfeng farm, W. P. Leu et al. 463 (HAST); Mt. Hohuanshan, Wuling to Kunyang, C. I Peng 8278 (HAST); Chingying Village to Yunhai, S. T. Chiu et al. 3332 (HAST); Tungpu Hot Spring to Rainbow Waterfall, C. I Peng 6475 (HAST); Chuntashan, T. Y. Liu et al. 956 (HAST); Prov. #21 131 km, T. Y. Liu et al. 1230 (HAST); Hsitou, M. T. Kao et al. 3166 (TAI); Fenghuangshan, C. I Peng et al. 17961 (HAST). Chiayi Co.: Peitzutung logging track, between Shihchuo and Tatungshan, C. C. Liao et al. 1678 (TAI); Fenchihu, S. F. Huang 2611 (TAI); Shihcho, Lungyun Lodge, K. Y. Wang et al. 736 (TAI); Alishan, County #169 Hwy, 44 km, S. M. Liu et al. 653 (TAI); Alishan, Prov. #18 Hwy 23-23.5 km, H. L. Ho et al. 1134 (TAI); Alishan Sacred Tree, T. Y. A. Yang et al. 6516 (TAI, TNM); Alishan to Chushan, T. Y. A. Yang et al. 7799 (TAI, TNM); Alishan, S. Sasaki et al. s.n. 17 May 1932 (TAI); Paiyun Lodge to Yushan main peak, C. I Peng et al. 14393 (HAST); Mt. Morrison, T. I. Chuang et al. s.n. 25 May 1960 (TAI). Tainan Co.: Kuantzuling, C. H. Chen et al. 1 (HAST). Kaohsiung Co.: Neimen, Shihkou, T. Y. A. Yang et al. 14476 (HAST); Liukuei, Y. Y. Huang et al. 236 (HAST); Meilan logging track 22-24 km, T. Y. Liu et al. 488 (HAST); Sanping, C. M. Wang et al. 1288 (HAST); Kuanshan, C. K. Liou 402 (HAST); Tienchi, C. H. Lin et al. 429 (HAST). Pingtung Co.: Kaoshihfo, G. P. Hsieh 2517 (PPI); Lilungshan, S. H. Wu et al. 1042 (HAST); Taiwu, Wanan, S. M. Ku 1210 (PPI); Laiyi, S. C. Wu 1213 (HAST); Wutai, Tawu village, S. H. Wu et al. 1155 (HAST); Tawushan, C. I Peng 13184 (HAST); Kuaiku Lodge to Mt. Peitawu, C. I Huang et al. 981 (HAST). Hualien Co.: Hsiaochingshui, S. Y. Lu 18842 (TAIF); Wanjung logging track 42.5 km, S. H. Wu et al. 1368 (HAST); Wanjung logging track to Chitsaihu, S. H. Wu et al. 1344 (HAST); Tungmen Check Point to Chilai electric power maitaining station, P. J. Lin 204 (HAST); Yuli, S. Y. Lu 1915 (TAIF). Taitung Co.: Chupang, M. T. Kao et al. 14234 (TAI); Taimali, Hsinhsing, Y. K. Chen 485 (HAST); Yenping logging track, C. F. Chen 1891 (PPI); Hsiangyangshan; S. T. Chiu et al. 2932 (HAST).
Note: Individuals of R. croceacanthus have variable morphologies, including shape and size of leaflets, serration of the margin, density of indumentum, and length of glandular hairs, features that probably reflect the diverse habitats they are able to grow from low to high altitudes. The taxonomic status of this species is very confusing due to many names proposed by Hayata (1918) and Koidzumi (1930) without detailed comparisons to the existing names. This confusion is further aggravated by misidentification of specimens and miscitation of names by the following researchers. We re-examined the isotype of R. croceacanthus var. glaber [Bunkiko, U. Faurie 46 (HAST)] and considered it to be a synonym of R.
croceacanthus here. Rubus rubro-angustifolius, treated as
a synonym of R. asper var. glaber by Hsieh (1989) and R.
croceacanthus var. glaber by Hsieh and Ohashi (1993), is
treated as a synonym of R. cardotii (see comments under the species description). One of the specimens [Mt. Chingsui, T. Shimizu et al. 11722 (TAI)] under the name
R. croceacanthus var. glaber (Hsieh and Ohashi, 1993)
should belong to R. linearifoliolus (see commnets under the species description).
An examination of the seven specimens cited by Hsieh (1989) under R. sumatranus revealed that, except S. F.
Huang 555 (TAI), others represent R. croceacanthus. The
can be distinguished by its larger flowers (2.5-5 cm in diam.), suborbicular petals, and globose to oblong and red aggregate fruits. Rubus sumatranus, on the contrary, has smaller flowers (1-2 cm in diam.), oblong petals, oblong and orange aggregate fruits. In addition, R.
dolichocephalus has been usually treated as a synonym
of R. sumatranus by several authers because of its ellipsoid aggregate fruits, but actually is closer to R.
croceacanthus and should be treated as synonym of the
latter because of its larger flowers and suborbicular petals.
The type specimens of R. hirsutopungens and R.
parvipungens are similar to each other; both names also
had been miscited to refer R. pungens var. oldhamii and
R. pungens by Hsieh and Ohashi (1993). Both of the two
taxa have rather small leaves and thinly glandular twigs, more likely present an ecotype of R. croceacanthus in high altitudes since the leaves of R. croceacanthus populations in high mountains may range from small to large and have barely to prominently glandular hairs. Finally, the type specimen [Mt. Shichiseitonzan, S.
Sasaki s.n. 1911 (TI)] of R. glanduloso-calycinus
[synonym of R. sumatranus in Hsieh (1989) and Hsieh & Ohashi (1993)] also have smaller and only slightly pubescent leaves, is treated as synonym of R.
croceacanthus here. Taken together, R. croceacanthus
exhibits tremendous variations in Taiwan and more indepth investigations are needed for this complex taxon in the future.
7. Rubus flagelliflorus Focke, Bot. Jahrb. Syst. 29: 393. 1900; Chang, Forest. J. Taiwan Prov. Pingtung. Inst.
Agr. 21: 1. 1979. 裂緣苞懸鉤子
Shrubs scandent or trailing. Twigs densely gray tomentose when young, glabrate in age, with sparse, curved minute prickles. Leaves simple; blade ovate to ovate-oblong, 7-9 × 5-7 cm, palmately 5-veined, lateral veins 4 or 5 pairs, raised abaxially, abaxially densely rusty yellow tomentose, adaxially hirsute along veins, base deeply cordate, margin often slightly undulate or undivided, irregularly serrate, apex acute to shortly acuminate; petiole 3-5 cm, densely gray tomentose when young, with sparse minute prickles; stipules caducous, free, to 9 mm, yellow tomentose-pilose, palmately lobed, lobes lanceolate. Inflorescences terminal, short racemes, 2.5-5 cm, few flowered, or flowers several in clusters; pedicel 1-2 cm, densely yellow tomentose-pilose; bracts to 10 mm, brown tomentose, palmately divided apically, lobes lanceolate. Flowers 1-1.2 cm in diam., hypanthium cup shaped; sepals reflexed after anthesis, ovate-lanceolate, 6-8 mm, abaxially densely yellow tomentose, outer sepal margin 2- or 3-lobed, inner sepals with entire margin, apex acuminate; petals white, 5-6 ×
4-5 mm, suborbicular, shorter than sepals; stamens many, glabrous or slightly pilose, anthers glabrous; carpels slightly longer than stamens, styles and ovaries glabrous. Aggregate fruit purplish black at maturity, semiglobose, 0.8-1 cm in diam., glabrous. Flowering season August to Octobor; fruiting season Octobor to December.
Distributed in Mainland China. Taiwan, found in Lanyu and Lutao; 200-500 m.
Specimens examined: TAIWAN: Taitung Co.: Lanyu, Hongtoushan, Y. L. Chun 553, 591 (PPI); W. C. Leong et al. 2484 (HAST); T. Y. A. Yang et al. 8101 (TNM); Lutao, C. E. Chang s.n. 30 Sep. 1982 (PPI).
8. Rubus formosensis Kuntze, Meth. Sp.-Beschr. Rubus,
73. 1879. 臺灣懸鉤子
Rubus rugosus Maxim., Bull. Acad. Imp. Sci. St-Péters. 17: 149. 1872, non Seem 1815.
Rubus formosanus Maxim. ex Focke, Biblioth. Bot. 17 (Heft 72): 117. 1910.
Rubus nantoensis Hayata, J. Coll. Sci. Univ. Tokyo 30: 92. 1911. Rubus randaiensis Hayata, J. Coll. Sci. Univ. Tokyo 30: 93. 1911.
Rubus rugosissimus Hayata, Icon. Pl. Formos. 3: 93. 1913.
Shrubs suberect or scandent. Twigs with soft yellowish gray tomentose hairs, unarmed or minutely prickled. Leaves simple; blade broadly ovate to suborbicular, 6-15 × 5-13 cm, 5-veined, reticulate veins raised and prominent abaxially, bullate between veins adaxially, abaxially yellowish gray tomentose, soft hairy along veins, adaxially soft hairy when young, base cordate, margin 3-5-lobed, lobes ovate-triangular, terminal one largest, margin irregularly coarsely serrate, apex acute or obtuse; petiole 2-4.5 cm, densely yellowish gray tomentose, usually unarmed; stipules caducous, free, brownish, broadly ovate to oblong, to 1.5 cm, entire or laciniate at apex, lobes lanceolate. Inflorescences terminal, short subracemes, 3-6 cm, several flowered, or flowers solitary in leaf axils; pedicel 3-5 mm, densely yellowish gray tomentose; bracts reddish brown, similar to stipules. Flowers 1-1.5 cm in diam., hypanthium cup shaped; sepals triangular to ovate, 5-8 mm, abaxially densely yellowish gray tomentose, margin entire or outer sepals apically with few teeth, apex acute to shortly acuminate; petals white, broadly suborbicular, 5-7 × 4-6 mm, base clawed, margin sometimes erose; stamens many, shorter than sepals, filaments linear, with few long hairs at apex of anthers; carpels slightly longer than stamens. Aggregate fruit yellow, orange-red or red at maturity, globose or broadly ovoid, 1-1.5 cm in diam., glabrous. Flowering season July to September; fruiting season September to November. 2n = 28.
Distributed in Southern China. Taiwan, commonly distributed in open slopes from medium to high altitudes, 500-3000 m throughout the central mountains.
Specimens examined: TAIWAN: Ilan Co.: Suao, Chimaoshan, T. Nakamura 5186 (TAI); Buta to Riyohen, T. Suzuki 7562 (TAI); Ssuchi to Liumaoan, S. Suzuki 4927 (TAI); Mimodeng, W. C. Leong et al. 3794 (HAST); Nanshan to Matang, C. H. Chen et al. 210 (HAST). Taoyuan Co.: Yangmei, Puhsin, W. C. Leong et al. 2728 (HAST); Shihmenshan, S. C. Liu et al. 712 (HAST); Tungyenshan, W. C. Huang 215 (HAST); Kappanzan, T. Suziki 4655 (TAI); Daman to Szleng, T. T. Chen 9234 (TAIF). Hsinchu Co.: Yuanyanghu, C. H. Chen et al. 180 (HAST); Leshan to Kuanwu, J. C. Wang et al. 4723 (TAI). Miaoli Co.: Shishihshan tunnel to Mt. Huoshih, C. M. Wang 1742 (HAST). Taichung City: Takeng, J. Y. Huang 1017 (HAST). Taichung Co.: Chingshan, S. F. Huang 551 (TAI); Huanshan, S. W. Chung 49 (HAST); Pachsienshan, M. T. Kao 2746 (TAI); Tahsuehshan Logging Station, T. S. Liu et al. 250 (TAI); Anmashan, C. M. Kuo 7103 (TAI); Tahsuehshan #230 logging tract, J. C. Wang et al. 4982 (TAI). Nantou Co.: Kuantaoshi, C. F. Hsieh et al. 1468 (TAI); Huisun Experimental Forest, S. M. Liu et al. 20 (HAST); Lienhuachi, K. Mori s.n. 8 Jul. 1936 (TAI); Meifeng, C. I Peng et al. 17172 (HAST); Chunda logging track 21 km, M. H. Chen et al. 26 (HAST); Kuankao to Tungpu, Y. F. Chen 3541 (TAI); Hsitou to Shanlinhsi, Y. F. Chen 2834 (TAI). Chiayi Co.: Chushan, Mori et al. s.n. 6 Nov. 1932 (TAI); Yushan, Masamune et al. s.n. 2 Jul. 1931 (TAI). Kaohsiung Co.: Taoyuan, Likuan to Tienchi, C. H. Lin et al. 416 (HAST); Liukuei to Peinanchushan, T. Yamazaki 399 (TAI); Shihshan, T. C. Huang 6526 (TAI); Prov. #20 119 km, C. C. Wang et al. 1041 (HAST). Pingtung Co.: Wutai to Ali, W. P. Leu et al. 997 (HAST); Ali, H. N. Yang 2022 (TAI); Chihpenchushan, Y. R. Lin et al. 218 (HAST); Chingshuiyin, H. Shimizu 3822 (TAI); Tahan logging track 17 km, K. F. Chung et al. 1267 (HAST). Hualien Co.: Nanan to Walami, C. C. Liao et al. 1546 (HAST); Chingshuishan, Shakatang logging track, C. C. Liao 478 (HAST); Tailukotashan, T. Suzuki 9287 (TAI); Mukuashan, T. Nakamura 4576 (TAI).
9. Rubus fraxinifolius Poir. in Lamarck, Encycl. Meth. 6: 242. 1806; Matsum. & Hayata, J. Coll. Sci. Univ.
Tokyo 22: 121. 1906. 榿葉懸鉤子
Rubus alnifoliolatus H. Lév., Bull. Soc. Bot. France 53: 549. 1906.
Rubus fraxinifolius var. kotoensis (Hayata) Koidz., J. Coll. Sci. Univ. Tokyo 34: 144. 1913. - Rubus kotoensis Hayata, Icon. Pl. Formos. 3: 90. 1913. - Rubus alnifoliolatus var. kotoensis (Hayata) Li, Woody Fl. Taiwan 320. 1963.
Shrubs suberect. Twigs dark-reddish, glabrous, sparsely prickled or sometimes unarmed, prickles curved, to 5 mm. Leaves imparipinnate, 7-9-foliolate, rarely 5-foliolate; blade of leaflets ovate-lanceolate to oblong-lanceolate, 5-10 × 1.5-3.5 cm, lateral veins 10-13 per side, raised abaxially and impressed adaxially, both surfaces glabrous or abaxially slightly pubescent along veins, base obtuse to rounded, margin irregularly serrate or doubly serrate, apex acute to acuminate, lateral leaflets often oblique; petiole reddish, 4-8 cm, glabrous, with sparse prickles; stipules attached to the petiole base, linear-lanceolate, to 10 mm, glabrous. Inflorescences terminal, usually broad cymose panicles, to 10 cm, many flowered; pedicel 1-1.4 cm, glabrous; bracts linear, to 9 mm. Flowers 1-1.5 cm in diam., hypanthium saucer shaped; sepals ovate- lanceolate, 7-11 mm, abaxially glabrous, with few sessile glands, apex acuminate to
caudate; petals white, ovate-oblong, 0.7-1.4 cm × 4-6 mm, abaxially with few hairs, apex obtuse; stamens many, glabrous, shorter than petals; carpels numerous, style and ovary glabrous, torus shortly stalked. Aggregate fruit red at maturity, oblong or ellipsoid, 1-1.5 × 0.8-1 cm. Flowering and fruiting season all year around. 2n = 14.
Distributed in the Philippines, Malaysia, Indonesia, and Pacific Islands. Taiwan, commonly distributed from low to medium altitudes, 20-2000 m throughout the island.
Specimens examined: TAIWAN: Taipei City: Chanchushan, T. Nakamura 2933 (TAI); Sanhsia, S. Sasaki et al. s.n. 25 Nov. 1910 (TAI); County #101A Hwy, near Yuyujen's grave, S. H. Wu 1444 (HAST). Taipei Co.: Manyuehyuan Waterfall, C. C. Liao et al. 378 (HAST); Pinglin, Prov. #9 Hwy 54 km; Hsintien, Kuanghsing, S. C. Liu et al. 618 (HAST); Lussutan, Y. C. Jeng 515 (TAI); Tonghouhsi, C. M. Kuo 9213 (TAI); Ayi, Wulai, M. T. Kao 30006 (TAI). Ilan Co.: Nanao, along Nanaonanhis, T. Y. Liu et al. 853 (HAST); Jentse, C. M. Wang 2358 (HAST); Tuchang to Ssuchi, S. Suzuki 14 (TAI); Nanshan to Matang, C. H. Chen et al. 238 (HAST); Ssuyuanyakou, T. Yamazaki et al. 1065 (TAI). Taoyuan Co.: Hsiayun to Hsiayunping, C. I Peng et al. 10298 (HAST); Fuhsing, Ishan village, C. C. Liao et al. 163 (HAST). Hsinchu Co.: Hsiatienpu, C. H. Yu 168 (HAST); Chenhsipao, S. C. Wu et al. 1827 (HAST); Tuchang to Kuanwu, C. I Peng 9371 (HAST). Miaoli Co.: Komokwan, Y. Shimada 863A (HAST). Taichung Co.: Wushihkeng, H. Y. Shen et al. 222 (HAST); Chiapaotai, J. P. Wu s.n. 13 Dec. 1981 (TAI); Techi, C. I Peng et al. 17068 (HAST). Nantou Co.: Puli, Kuanyin waterfall, C. L. Wu s.n. 29 Jul. 1970 (TAI); Lushan, C. I Peng 6149 (HAST); Lihsing Private Rd., W. C. Leong et al. 1277 (HAST); Hsinyi, Lona, C. I Peng et al. 17001 (HAST); Hoshe, T. C. Huang et al. 15485 (TAI). Chiayi Co.: Alishan, Tefuyeh, W. C. Leong et al. 2561 (HAST). Kaohsiung Co.: Tengchih lodging area to Shihshan, Y. R. Lin 442 (HAST); Tengchih, T. C. Huang 15897 (TAI); Shanping, T. Kiang et al. 91 (TAI). Pingtung Co.: Wutai to Ali, W. P. Leu et al. 995 (HAST); Ali to Hsiaokueihu, C. C. Liao et al. 576 (HAST); Chingshuiyin, H. Shimizu 3614 (TAI); Tahan logging track 13-14 km, K. F. Chung et al. 1222 (HAST); Mt. Wanlite, T. Y. A. Yang et al. 2755 (HAST); Shouka, M. T. Kao et al. 3491 (TAI). Hualien Co.: Hoping logging track 15 km, S. M. Liu et al. 357 (HAST); Tailuko, M. T. Kao 4093 (TAI); Honku, M. T. Kao 9839 (TAI); Tienhsian, M. T. Kao et al. 4416 (TAI); Hungyeh Hot Spring, C. I Peng et al. 11599 (HAST); Wanjung logging track 23.8 km, S. H. Wu et al. 1393 (HAST). Taitung Co.: Chengkung, Keng & Kao 2688 (TAI); Chinwha to Tahwu, T. S. Liu et al. 2861 (TAI); Taiwan Cycas Nature Reserve, S. M. Liu et al. 458 (HAST); Yenping logging track; S. M. Liu et al. 460 (HAST); Lichia logging track, Y. C. Kao et al. 152 (HAST); Darun Farm, Y. P. Cheng 3326 (TAIF); Kotosyo, S. Sasaki s.n. Jun. 1926 (TAI); Lanyu, M. T. Kao 5428, 8735 (TAI); Lanyu, Langtao, C. L. Yeh 95 (PPI); Lanyu, Between Hongtou Village and Tienchi, C. I Peng 5246 (HAST); Lutao, Ameishan, Hayashi 13 (TNM); Lutao, C. E. Chang 11713 (PPI).
Note: For a long time, the name R. alnifoliolatus had been used to refer to this taxon, while R. fraxinifolius had been used to refer to the unarmed population in Lanyu. We found the twigs in the isotype of R. fraxinifolius [Java,
Commerson s.n. (P)] are armed with sparse prickles, and
also observed the prickled and unarmed individuals in a same population in Lanyu. We treat R. alnifoliolatus as a synonym of R. fraxinifolius here according to nomenclature priority and also follow the note on HAST specimens by Naruhashi (2002).
10. Rubus howii Merr. & Chun, Sunyatsenia 5: 71. 1940; Lu & Yang, Taiwania 25: 123. f. 2. 1980.
裂葉懸鉤子 Shrubs straggling. Twigs densely yellowish brown pilose, with curved, minute prickles. Leaves simple; blade oblong to ovate-oblong, 4-8 × 2-5 cm, lateral veins 6-8 pairs, prominent abaxially and impressed adaxially, both surfaces yellowish brown pilose along veins, abaxially with sparse, minute prickles along midvein, base cordate, margin irregularly coarsely serrate, with 1 or 2 broadly ovate shallow lobes on each side below middle, apex acute to shortly acuminate; petiole 1-2 cm, densely yellowish brown pilose, with few small prickles; stipules caducous, free, 6-9 mm, deeply lobed, lobes linear, yellowish brown pilose. Inflorescences terminal, subracemes, 4-6 cm, few flowered, or flowers several in clusters in leaf axils; pedicel ca. 1 cm, densely yellowish brown pilose, with sparse prickles; bracts 5-8 mm, lobes linear, yellowish brown pilose. Flowers ca. 1 cm in diam., hypanthium cup shaped; sepals ovate-lanceolate, 6-8 mm, margin entire, apex acuminate; petals white, elliptic, 5-7 × 4-5 mm, base shortly clawed, apex obtuse; stamens many, shorter than petals, filaments glabrous; carpels glabrous, longer than stamens. Aggregate fruit red at maturity, globose, 1-1.5 cm in diam., glabrous. Flowering season March to May; fruiting season May to July.
Distributed in Hainan and Taiwan. Taiwan, rarely found in forests in the eastern part of the island at medium altitudes 800-1500 m.
Specimens examined: TAIWAN: Hualien Co.: Chinshui Farm, J. Y. Huang 2312 (HAST). Taitung Co.: Lichia logging tract, J. Y. Huang 2340 (HAST); S. Y. Lu 22113 (TAIF); Yenping forest road, Y. J. Lin 200 (PPI).
Note: This taxon in Hainan is rather rare and with very few collections, the information about fruit is absent in original literature and the Flora of China. Here is the first record about color and shape of aggregate fruit. The species was neglected in Hsieh (1989) and Hsieh & Ohashi (1993), and had no other collection since the first description. We newly found several populations at the medium altitudes in Hualien and Taitung County; a few specimens wrongly identified or unidentified in herbarium are also found.
11. Rubus incanus Sasaki ex T.-S. Liu & T.-Y. Yang, Quart. J. Taiwan Mus. 20: 167. 1968. 白絨懸鉤子
Rubus niveus auct. non Thunb.: Hsieh, J. Taiwan Mus. 42: 112. 1989.
Shrubs suberect or scandent. Twigs tomentose when young, glabrate in age, sparsely prickled. Leaves imparipinnate, 5-7-foliolate, sometimes 3-foliolate on flower bearing twigs; blade of leaflets ovate-elliptic to rhombic, midrib and lateral veins abaxially raised and minutely prickled, adaxially slightly impressed, abaxially gray tomentose, adaxially soft hairy along veins or glabrate, terminal leaflet subrhombic, slightly longer than lateral leaflets, 4-7 × 2-4 cm, base cuneate to rounded, margin double-serrate, becoming entire toward the base, apex acute to acuminate, lateral leaflets rhombic-elliptic, 2-4.5 × 1-2.5 cm, base oblique; petiole 1.5-4 cm, tomentose, with sparse prickles; stipules attached to the petiole base, linear, to 8 mm long, pubescent. Inflorescences terminal, narrow cymose panicles or short thyrses, 5-10 cm; pedicel 5-10 mm, tomentose; bracts lanceolate or linear, pubescent. Flowers to 1 cm in diam., hypanthium saucer shaped; sepals 4-6 mm, abaxially densely tomentose, with intermixed soft hairs, apex caudate or abruptly pointed; petals pink to purplish red, unopened, suborbicular, 4-6 × 3-5 mm, shorter than sepals, base clawed; stamens many, nearly as long as petals; carpels 65-90, slightly longer than stamens, ovary gray tomentose. Aggregate fruit red at maturity, globose, 0.8-1.2 cm in diam., hairy. Flowering season June to July; fruiting season August to September. 2n = 14.
Endemic to Taiwan. Commonly distributed in open places or forest edges at medium altitudes, 1800-3000 m throughout the central mountains.
Specimens examined: TAIWAN: Ilan Co.: Ssuyuanyakou, C. I Peng et al. 17409 (HAST). Hsinchu Co.: Talu logging track, J. C. Wang et al. 4864 (TAI); Kuanwu, Talu east branch, 3.4 km, L. J. Juan et al. 21 (HAST). Taichung Co.: Chingshan to Kukuan, T. C. Huang et al. 9629 (TAI); #710 logging track, J. C. Wang et al. 3591 (TAI); Yunleng Lodge, C. I Huang et al. 2612 (HAST); Nanhutashan, M. T. Kao 8617 (TAI); Chungyangchien, N. Fukuyama et al. 15206 (TAI). Nantou Co.: Tsuifeng, W. H. Hu 1432 (HAST); Juiyen Protection Area, C. H. Chen et al. 1257 (HAST); Pilushan logging track, J. J. Chen et al. 397 (HAST); Nengkaoshan, E. Matuda s.n. 4 Aug. 1919 (TAI). Kaohsiung Co.: Chinching Bridge, K. F. Chung et al. 1084 (HAST); Shishan logging track, G. P. Hsieh 629 (HAST, PPI). Hualien Co.: Hopin logging track, S. F. Huang et al. 5119 (TAI); Chilai, E. Matuda s.n. 4 Aug. 1919 (TAI); Mukuashan, M. T. Kao 4165 (TAI); Tarokotaizan, T. Suzuki 9613 (TAI); Pilushan to Pilushan Shenmu, C. I Peng 9254 (HAST). Taitung Co.: Hsiangyang, C. I Peng et al. 11829 (HAST).
Note: Rubus incanus was treated as a synonym of R.
niveus by Hsieh (1989). After comparing the populations
of Mainland China and Taiwan, we consider R. incanus and R. niveus are two distinct taxa. The inflorescences of
R. incanus are narrow cymose panicles or short thyrses,
12. Rubus inopertus (Focke ex Diels) Focke, Biblioth. Bot. 17 (Heft 72): 182. 1911; Naruhashi, Bot. Bull. Acad. Sin. 43: 194. 2002.
紅花懸鉤子（李棟山懸鉤子）
Rubus ritozanensis Sasaki, Trans. Nat. Hist. Soc. Formos. 21: 250. 1931.
Rubus fraxinifolius Poir. var. yushuni Suzuki & Yamam. Trans. Nat. Hist. Soc. Formos. 22: 409. 1932. - Rubus yushuni (Suzuki & Yamam.) Suzuki & Yamam., Trans. Nat. Hist. Soc. Formos. 25: 130. 1935.
Rubus ohwianus Koidz., Acta Phytotax. & Geobot. 8: 108. 1939.
Rubus yamamotoanus H.-L. Li, Woody Fl. Taiwan 319. 1963.
Shrubs suberect or scandent. Twigs glabrescent, with sparse prickles. Leaves imparipinnate, usually 9-11-foliolate; blade of leaflets ovate to ovate-lanceolate, 3-8 × 1.5-4 cm, lateral veins 9-12 per side, raised abaxially and impressed adaxially, both surfaces nearly glabrous, slightly pubescent along veins, base rounded to truncate, margin coarsely sharp doubly serrate, apex acuminate to caudate; petiole 3.5-5 cm, glabrous or slightly pubescent, with sparse, curved minute prickles; stipules attached to the petiole base, linear-lanceolate, to 5 mm, nearly glabrous. Inflorescences terminal and axillary, umbellate cymes, 2-4 cm, several flowered, or flowers several in axillary clusters; pedicel 4-8 mm, glabrous, unarmed; bracts linear-lanceolate, to 5 mm, nearly glabrous. Flowers 1-1.2 cm in diam., hypanthium saucer shaped; sepals reflexed in fruit, triangular-ovate, 4-6 mm, glabrous, inner sepals tomentose at margin, apex acute to acuminate; petals pink to purplish red, unopened, obovate, 5-7 × 4-6 mm, base slightly hairy and shortly clawed; stamens numerous, slightly shorter than petals; carpels 65-80, slightly longer than stamens, ovary and base of style soft hairy. Aggregate fruit dark red to dark purple at maturity, globose, 0.6-1 cm in diam., thinly pubescent. Flowering season May to June; fruiting season July to August. 2n = 14.
Distributed in Mainland China and Vietnam. Taiwan, occurred in moist forests edges in the northern and central parts at medium altitudes 1500-2300 m.
Specimens examined: TAIWAN: Ilan Co.: Chiaping logging track, near a waterfall, C. H. Lin et al. 594 (HAST); Taipingshan, C. I Peng et al. 7879 (HAST); Chiliting, T. C. Huang 7826 (TAI); Siyuanyakou, W. C. Leong et al. 3773 (HAST). Taoyuan Co.: Lalashan, S. Y. Lu 22221 (TAIF). Hsinchu Co.: Yuanyang Lake, #170 logging track 5.5 km, W. C. Leong et al. 2364 (HAST); Litung Lodge, Y. Y. Huang et al. 448 (HAST); Kuanwu, Shenmu logging track, L. J. Juan et al. 36 (HAST). Taichung Co.: #710 logging track, W. C. Leong et al. 2401 (HAST). Nantou Co.: Mayfeng, water source area, C. I Peng 9113 (HAST); Juiyenhsi Forest Reserve, J. J. Chen et al. 420 (HAST). Chiayi Co.: Alishan, C. E. Chang 3910 (PPI). Hualien Co.: Hopin logging track, S. F. Huang et al. 5117 (TAI).
Note: The treatment is based the note Naruhashi wrote on the specimens of R. ritozanensis in HAST in 2002 or earlier and we also agree this point of view.
13. Rubus lambertianus Ser. in Candolle, Prodr. 2: 567. 1825; Matsum. & Hayata, J. Coll. Sci. Univ. Tokyo 22: 121. 1906.
13a. Rubus lambertianus var. lambertianus 高粱泡
Rubus lambertianus Ser. ssp. xanthoneurus auct. non Focke: Koidz., J. Coll. Sci. Univ. Tokyo 34: 158. 1913.
Shrubs straggling. Twigs brown-yellow velutinous, especially when young, with sparse, curved prickles. Leaves simple; blade broadly ovate, 4-12 × 3-8 cm, 5-veined, both surfaces pilose, more densely so along veins, with sparse, minute prickles along midvein, base broadly cordate to rounded, margin indistinctly 3-5-lobed or undulate, serrulate or dentate, apex acuminate; petiole 2-4 cm, thinly pubescent, with sparse, minute prickles; stipules caducous, free, to 1 cm, shallowly to deeply laciniate, lobes linear to lanceolate, thinly pubescent. Inflorescences terminal, broad cymose panicles, 10-20 cm, sometimes flowers few in subracemes or clusters in leaf axils; pedicel 8-12 mm, thinly pubescent; bracts to 5 mm, similar to stipules. Flowers 7-9 mm in diam., hypanthium cup shaped; sepals triangular-ovate, 4-7 mm, abaxially thinly pubescent, apex acuminate, sometimes with few teeth; petals white, elliptic to obovate, 3-6 × 2-5 mm, slightly shorter than sepals, base slightly hairy and clawed; stamens many, somewhat shorter than petals; carpels ca. 15-20, shorter than stamens, glabrous. Aggregate fruit orange-red or yellow at maturity, globose, 0.6-0.9 cm in diam., glabrous. Flowering season August to October; fruiting season October to December. 2n = 28.
Distributed in Mainland China and Japan. Taiwan, distributed in thickets from low to medium altitudes, 100-2000 m throughout the island.
Specimens examined: TAIWAN: Ilan Co.: Chiaping logging track, C. H. Lin et al. 612 (HAST); Mt. Taiping, C. C. Chuang et al. 4780 (HAST); Nanshan to Matang, C. H. Chen et al. 228 (HAST). Taoyuan Co.: Paling to Kaoyi, C. I Peng 11052 (HAST); Fuhsing, Ronghuaciao, C. C. Chen 511 (HAST). Hsinchu Co.: Hengshan, Dashanbeishan, S. C. Wu 2424 (HAST); Wuchihshan, H. F. Yen 8902 (TAI); Chienshih, Meihua Shechu, C. H. Lin 499 (HAST); Litung Lodge, C. M. Wang et al. 1984 (HAST). Miaoli Co.: Nanchuang, Fengmeihsi bank, T. Y. Liu et al. 361 (HAST); Henglongkutao, C. H. Yu 404 (HAST). Taichung City: Takeng, C. H. Ou et al. s.n. 17 May 1990 (TNM). Taichung Co.: Tungshih, #200 logging tract 8 km, J. Y. Huang 2326 (HAST); Wushihkeng, S. W. Chung 3517 (TAIF); Kuangming Bridge to Techi Reservoir, W. P. Leu et al. 360 (HAST); Lishan: Prov. #8 Hwy 54 km, T. Y. Liu et al. 292 (HAST). Nantou Co.: Hsinyi, Lona, C. I Peng et al. 16995 (HAST); Hsini to Hoshe, W. P. Leu et al. 1074 (HAST);
